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Home , Azolla filiculoides, Azolla pinnata, Azolla rubra, Salvinia, Salviniaceae, Salviniales

FLORA OF AND LYCOPHYTES

SALVINIACEAE

P.J. BROWNSEY & L.R. PERRIE

Fascicle 10 – MARCH 2015 © Landcare Research New Zealand Limited 2015. Unless indicated otherwise for specific items, this copyright work is licensed under the Creative Commons Attribution 3.0 New Zealand license. Attribution if redistributing to the public without adaptation: “Source: Landcare Research" Attribution if making an adaptation or derivative work: “Sourced from Landcare Research" See Image Information for copyright and licence details for images.

CATALOGUING IN PUBLICATION Brownsey, P.J. (Patrick John), 1948- Flora of New Zealand [electronic resource] : ferns and lycophytes. Fascicle 10, / P.J. Brownsey and L.R. Perrie. -- Lincoln, N.Z. : Manaaki Whenua Press, 2015. 1 online resource ISBN 978-0-478-34779-1 (pdf) ISBN 978-0-478-34761-6 (set) 1.Ferns -- New Zealand - Identification. I. Perrie, L.R. (Leon Richard). II. Title. III. Manaaki Whenua- Landcare Research New Zealand Ltd. UDC 582.394.771(931)DC 587.30993

DOI: 10.7931/B1H59T

This work should be cited as: Brownsey, P.J. & Perrie, L.R. 2015: Salviniaceae. In: Breitwieser, I.; Heenan, P.B.; Wilton, A.D. Flora of New Zealand — Ferns and Lycophytes. Fascicle 10. Manaaki Whenua Press, Lincoln. http://dx.doi.org/10.7931/B1H59T

Cover image: rubra, floating on the surface of standing water with irregularly branched stems bearing densely imbricate bilobed floating leaves Contents

Introduction...... 1 Taxa Salviniaceae Martinov ...... 2 Azolla Lam...... 2 R.Br...... 3 R.Br...... 5 Ség...... 6 Salvinia ×molesta D.S.Mitch...... 7 References ...... 10 Acknowledgements ...... 12 Maps ...... 13 Index ...... 15 Image Information ...... 16

Introduction

The Salviniaceae is represented in New Zealand by two genera, Azolla and Salvinia. Azolla has one indigenous and one naturalised , whilst Salvinia is represented by a single species which has been declared a noxious weed. Azolla rubra is indigenous, occurring in slow-moving streams and ponds throughout the North Island and the eastern half of the South Island. Azolla pinnata was first recorded in 1969, and is largely confined to the northern North Island, but is now spreading southwards, and has replaced A. rubra in some parts of Northland. Salvinia ×molesta is an extremely aggressive weed of lakes and ponds in Northland and Auckland. Both genera are free-floating, aquatic ferns with highly modified leaves adapted for flotation; they are characteristically heterosporous, producing mega- and microsporangia in hardened sporocarps. Azolla has small leaves and forms red carpets of growth on the water surface. Salvinia has dimorphic leaves arranged in whorls of three, and in S. ×molesta, characteristic egg-whisk-shaped hairs on the floating leaves for buoyancy.

1 Salviniaceae Martinov, Tekhno-Bot. Slovar, 559 (1820) = Azollaceae Wettst., Handb. Syst. Bot. 2, 77 (1903)

Type taxon: Salvinia Ség. Aquatic ferns, free-floating. Stems slender, branching, glabrous or hairy; roots either long and pendent, or absent. Leaves either monomorphic and bilobed, with the upper lobe aerial and green and the lower colourless and floating, or dimorphic and arranged in whorls of 3, with two floating and one submerged; not articulated, herbaceous, papillate or with very specialised hairs, non-circinate when young. Veins anastomosing, or absent. Sporocarps globose or conical, sessile or stalked, glabrous or hairy, either attached to the lower lobes of bilobed leaves, or to the submerged leaves, splitting open irregularly, each bearing either mega- or microsporangiate sori. Annulus absent. Heterosporous, trilete, lacking chlorophyll; one in each megasporangium and 32–64 in each microsporangium. spheroidal, plain to tuberculate; microspores spheroidal, and plain to rugulate. : A family of two genera and about 17 species. The Salviniaceae comprises floating aquatic ferns with highly modified leaves adapted for flotation, mega- and micro-sporangia borne in hardened sporocarps that give rise to mega- and micro-spores, and spores which germinate into greatly reduced still attached to the parent . Pryer (1999) and Pryer et al. (2004) showed that the heterosporous ferns, , are one of three major lineages of “core leptosporangiate” ferns, along with tree ferns and polypod ferns. Nagalingum et al. (2008) subsequently confirmed that Salviniales is divided into two strongly supported families, the and Salviniaceae, and that within Salviniaceae there is strong support for recognition of two genera, Azolla and Salvinia. Schneider & Pryer (2002) also demonstrated that the spores of Marsileaceae are markedly different to those of Salviniaceae and all homosporous ferns. The morphology of the reproductive structures in heterosporous water ferns has been critically examined by Nagalingum et al. (2006). Some authors (Schneller 1990a; Chinnock 1998; Jones 1998) recognise two separate families, Azollaceae and Salviniaceae, on the basis of the significant differences in the morphology of the two groups, but we follow Smith et al. (2006) and Christenhusz et al. (2011) in recognising just one family. It is represented in New Zealand by one indigenous (Azolla) and one fully naturalised genus (Salvinia). Distribution: Widespread in tropical and temperate regions. One native and one naturalised genus with three species in New Zealand; none endemic. Biostatus: Indigenous (Non-endemic). Table 1: Number of species and named hybrids in New Zealand within Salviniaceae Martinov Category Number Indigenous (Non-endemic) 1 Exotic: Fully Naturalised 2 Total 3 Azolla Lam., Encycl. 1, 343 (1783) Type taxon: Lam. Etymology: From the Greek azo (to dry), and ollyo (to kill), a reference to the fact that plants are killed by drying out. Aquatic ferns, free-floating. Stems short, branching, glabrous or sometimes hairy, bearing pendent roots. Leaves monomorphic, bilobed; the upper lobe aerial, green, papillate, containing the cyanobacterium in a large cavity on the lower surface; the lower lobe colourless and floating; both lobes glabrous. Veins absent. Sporocarps usually absent in NZ; when present, borne on the first leaf of a lateral branch, in the axil of the lower lobe and protected by the upper lobe; megasporocarps oblong-conical; microsporocarps ovoid to globular, usually larger than megasporocarps. Megaspores solitary in each megasporangium, bearing hair-like filaments at the base and floats (massulae) at the apex. Microspores 32–64 in each microsporangium, mixed with massulae that bear hooked processes (glochidia). Megaspores spheroidal, coarsely tuberculate, perforate; microspores spheroidal, plain.

2 1 Plants regularly 2-3-pinnate; roots with fine lateral rootlets ...... pinnata Plants irregularly branched; roots simple ...... rubra Distribution: Seven species widespread in tropical and temperate regions (Reid et al. 2006); four occur in North and , two in and , and two in , one of which is also native to New Zealand and the other introduced. One naturalised and one native species in New Zealand. Biostatus: Indigenous (Non-endemic). Table 2: Number of species in New Zealand within Azolla Lam. Category Number Indigenous (Non-endemic) 1 Exotic: Fully Naturalised 1 Total 2 Cytology: The base chromosome number in Azolla is x = 22 (Schneller 1990a). Notes: Plants of Azolla are free-floating and capable of covering large areas of slow-moving or still water. They are easily recognised as a red carpet of growth on the water surface. In New Zealand, fertile plants are rarely found, and reproduction is usually vegetative by fragmentation of the lower branches of the plants. Only five collections with sporocarps have been detected in New Zealand herbaria (AK 284894, CHR 168181, 214754, WELT P002240, P024125). Azolla is of considerable economic importance in Asian countries where crops such as rice are grown in flooded fields. Azolla contains the cyanobacterium Anabaena which fixes and is a source of fertiliser when the parent dies. The phylogenetic relationships of all seven species of Azolla have been investigated by Reid et al. (2006) and Metzgar et al. (2007). Spores are illustrated by Tryon & Lugardon (1991). Azolla pinnata R.Br., Prodr. Fl. Nov. Holland., 167 (1810) Lectotype (selected by Brownsey & Perrie 2014): Patersons River [New South Wales], R. Brown Iter Austral. No. 135, Oct. 1804, BM 001048353!

Etymology: From the Latin pinnatus (pinnate), a reference to the regular pinnate branching of the plant. Vernacular name: ferny azolla Aquatic , free-floating, forming extensive red or green mats. Plants usually broadly triangular or pentagonal or occasionally ovate or elliptic, 5–25 mm long, 4–25 mm wide; stems branching regularly, usually 2-pinnate, or sometimes 3-pinnate in the distal half of the lower pinnae; roots with fine lateral rootlets, 10–60 mm long. Leaves densely imbricate; upper lobe of leaves ovate to elliptic, 0.5–1.5 mm long, 0.3–0.8 mm wide, red or green, papillate; apices acute to obtuse; margins membranous and translucent. Sporocarps rarely produced; megasporocarps not seen; microsporocarps ovoid to spheroidal, covered in a membranous indusium, 1.0–2.0 mm long, 0.8–1.5 mm wide, apiculate, glabrous.

3 Distribution: North Island: Northland, Auckland, Volcanic Plateau, Taranaki, Southern North Island. Altitudinal range: 0–440 m. A widespread species of tropical and warm temperate Africa, Asia, Australia and New Caledonia, now naturalised in ponds and lakes throughout Northland, Auckland, Waikato and the Bay of Plenty, south to Kāwhia and Lake Ohakuri near Atiamuri. There are also specimens collected since 2009 from New Plymouth, Wanganui, Turakina, Otaki and Pekapeka, near Waikanae, suggesting that the plant is spreading southwards. It occurs from sea level up to 440 m at Lake Rerewhakaaitu in the Rotorua region. In Northland, it appears to be replacing the native A. rubra. Biostatus: Exotic; fully naturalised. Habitat: Grows on the surface of farm ponds, ornamental ponds, lakes, sluggish stream margins and in dune swamps. It often occurs with Spirodela, Wolffia, Lemna, Landoltia and Potamogeton cheesemanii, and occasionally with A. rubra, sometimes amongst Typha orientalis or sedges, or on open water. Fig. 1: Azolla pinnata distribution map based on databased records at AK, First record: NZ Weed & Pest Control Society (1969). Voucher: CHR and WELT. CHR 169486, 1969. Recognition: Azolla pinnata is readily distinguished from A. rubra by its more regular branching and by its roots with fine rootlets. It occurs only in the North Island. Notes: Reid et al. (2006) and Metzgar et al. (2007) showed that Azolla pinnata and the African A. nilotica form a sister clade to A. rubra, A. filiculoides and the other American species.

Fig. 2: Azolla pinnata: plants floating on the Fig. 3: Azolla pinnata: plant floating on the surface surface of standing water with regularly of standing water with regularly branched stems branched stems bearing bilobed floating leaves. bearing densely imbricate, ovate, papillate, floating leaves.

Fig. 4: Azolla pinnata: plants covering surface of standing water, tinged red in open sun.

4 Azolla rubra R.Br., Prodr. Fl. Nov. Holland., 167 (1810) ≡ Azolla filiculoides var. rubra (R.Br.) Strasb., Über Azolla, 78 (1873) Lectotype (selected by Brownsey & Perrie 2014): Budgery's Bridge [Badgery's Bridge, Nepean River, New South Wales], R. Brown, Apr. 1805, BM! (photo WELT E463/16)

Etymology: From the Latin ruber (red), a reference to the colour of the plant when growing in exposed habitats. Vernacular names: Pacific azolla; kārearea; kārerarera; retoreto; returetu Aquatic fern, free-floating, forming extensive red or green mats. Plants broadly ovate, elliptic or broader than long in outline, 6–30 mm long, 5–40 mm wide; stems irregularly branched; roots simple, 5–70 mm long. Leaves densely imbricate; upper lobe of leaves ovate, broadly ovate or broader than long, 0.5–2.0 mm long, 0.5–1.5 mm wide, red or green, papillate; apices obtuse to rounded; margins membranous and translucent. Sporocarps rarely produced; megasporocarps oblong-conical, c. 0.5 mm long; microsporocarps spheroidal, covered in a membranous indusium, 1.0–1.5 mm long, apiculate; both glabrous. Distribution: North Island: Northland, Auckland, Volcanic Plateau, Taranaki, Gisborne, Southern North Island. South Island: Western Nelson, Sounds-Nelson, Marlborough, Westland, Canterbury, Otago, Southland, Fiordland. Altitudinal range: 0–800 m. Azolla rubra occurs in scattered lowland areas of Northland, where it is being replaced by the naturalised A. pinnata. It is more common through Auckland, Waikato, the Bay of Plenty south to Lake Taupō, and in the southern North Island. On the east coast it has been collected as far north as Māhia, but is apparently absent from much of the East Cape district. It is also absent from much of Taranaki. It occurs most frequently in lowland areas up to c. 300 m but reaches almost 800 m in the Kaimanawa Mountains. In the South Island it is largely confined to lowland areas east of the main divide, extending westwards to near Karamea in the north, and Lake Manapouri in the far south. It extends from sea-level to c. 550 m near Lewis Pass. Also Australia (all States), and probably Papua New Guinea, Indonesia and (Reid et al. 2006). Fig. 5: Azolla rubra distribution map Biostatus: Indigenous (Non-endemic). based on databased records at AK, Habitat: Occurs on slow moving streams, ponds, lakes and swamps, CHR and WELT. and in farm ponds, ditches and drains where it forms extensive red mats of vegetation. It often occurs with species of Spirodela, Wolffia, Lemna, Landoltia, Ludwigia and Potamogeton, and occasionally with A. pinnata, sometimes amongst Typha orientalis, Carex secta or Salix species, or on open water. Recognition: Azolla rubra is readily distinguished from A. pinnata by its irregular branching and by its roots that lack fine rootlets. Azolla rubra is currently the only species that occurs in the South Island, but the southward spread of A. pinnata into the Wellington region suggests that it may soon cross Cook Strait. Cytology: No count has been made from New Zealand material of Azolla rubra, but 2n = 44 has been reported for Australian material (Tindale & Roy 2002). Notes: In New Zealand, this plant was treated as A. rubra by Allan (1961) and by all previous Flora writers. However, following Svenson (1944), the name A. filiculoides was adopted for the New Zealand plant by Brownsey et al. (1985) and Brownsey & Smith-Dodsworth (2000), and for the Australian plant by Chinnock (1998). In a detailed investigation of the genus, Reid et al. (2006) showed that A. filiculoides and A. rubra can be distinguished on both morphological characters and molecular differences. Metzgar et al. (2007) confirmed that the two are closely related sister taxa that are themselves sister to a clade containing the remaining American species. The name A. rubra was therefore reinstated by Brownsey & Perrie (2013) as a species native to Australia and New Zealand, and sister to the American A. filiculoides.

5 Fig. 6: Azolla rubra: plants floating on the Fig. 7: Azolla rubra: plants floating on the surface surface of standing water with irregularly of standing water showing the upper lobes of branched stems bearing densely imbricate bilobed floating leaves, tinged red. bilobed floating leaves

Fig. 8: Azolla rubra: simple roots hanging down Fig. 9: Azolla rubra: plants covering the surface of from plants floating on the water surface. standing water, tinged red in the open sun.

Salvinia Ség., Pl. Veron. 3, 52 (1754) Type taxon: (L.) All. Etymology: Named in honour of Professor Antonio Maria Salvini (1633-1722), botanist and Greek scholar from Florence. Aquatic ferns, free-floating. Stems short, branching, hairy, lacking roots. Leaves dimorphic, arranged in whorls of 3; each group comprising two floating, green, entire, foliar leaves, and one submerged, highly divided leaf appearing root-like; upper surface of foliar leaves bearing specialised water- repellent hairs; lower surface of foliar leaves and the submerged leaves bearing multicellular hairs. Veins anastomosing without free included veinlets. Sporocarps attached to the submerged leaves, subsessile or stalked, globose to ovoid; each sporocarp bearing either mega- or microsporangia. Sporangia usually empty in NZ plants; when fertile (not NZ), megaspores solitary in each megasporangium; microspores 32–64 in each microsporangium. Spores absent or aborted in NZ plants; when formed (not NZ) megaspores spheroidal, plain, perforate; microspores spheroidal, plain to rugulate. Distribution: 10–12 species widespread in tropical regions, extending also into temperate areas, with two centres of distribution in Africa, and in South and (Schneller 1990b); some species have become tropical weeds. One naturalised species in New Zealand. Biostatus: Exotic; fully naturalised.

6 Table 3: Number of species and named hybrids in New Zealand within Salvinia Ség. Category Number Exotic: Fully Naturalised 1 Total 1 Cytology: x = 9 (Schneller 1990b), the lowest base chromosome number known in ferns (Smith et al. 2006). Notes: The taxonomy of neotropical species was outlined by Mitchell & Thomas (1972) and adopted by Tryon & Tryon (1982). Following the initial investigation, Salvinia ×molesta was described as a new species (Mitchell 1972). It belongs to a small group of South American taxa, including S. auriculata and S. biloba (syn. S. herzogii, see de la Sota 1995), in which the hairs at the end of the leaf papillae are joined at their distal ends. Based on analysis of five different genes, Nagalingum et al. (2008) showed that Salvinia comprises two Eurasian and American groups, although only the American clade, which includes S. ×molesta, is strongly supported. Plants of Salvinia are free-floating and capable of covering large areas of slow-moving or still water. In New Zealand, sporocarps are rare and are usually empty when found, and reproduction is vegetative. Spores of American species are illustrated by Tryon & Lugardon (1991). Salvinia ×molesta D.S.Mitch., Brit. Fern Gaz. 10: 251 (1972) Holotype: Ruziruhuru River inlet, Lake Kariba, Rhodesia [Zimbabwe], D.S. Mitchell 1330, SRGH.

Etymology: From the Latin molestus (troublesome), a reference to its weedy and invasive nature. Vernacular names: Kariba weed; salvinia Aquatic fern, forming free-floating dense mats, with leaves tightly overlapping. Floating leaves broadly elliptic to orbicular when young, becoming broader than long at maturity, 12–30 mm long, 15–50 mm wide, often conduplicately folded along the midrib; apex emarginate; margins entire, sometimes inrolled; base cordate, shortly stalked; upper surface light or brownish-green, becoming darker on the margins, densely covered in papillae each bearing 2 or 4 uniseriate hairs united at their distal ends (shaped like an egg-whisk); papillae up to 3 mm long. Submerged leaves branched, 25–300 mm long, densely covered in brown septate hairs 1–3 mm long. Fertile axes on submerged leaves 15–70 mm long, bearing 4–20 pairs of sporocarps arranged in 2 rows. Sporocarps spheroidal or ovoid, 2–3 mm long, 1–2 mm wide, densely hairy, subsessile, containing sporangia that are usually empty or produce only aborted spores. Distribution: North Island: Northland, Auckland. Altitudinal range: 0–50 m. A rampant weed of tropical and warm temperate regions, recorded as a naturalised plant in lowland areas of northern New Zealand from near Kaitaia to Hamilton. There are additional records of the plant from garden ponds as far east and south as Wairoa (AK 245013). Biostatus: Exotic; fully naturalised. Habitat: Recorded as an aquatic weed on lakes and ornamental ponds, and in Eleocharis sphacelata swamp. First record: Mason (1964, p. 235), Armiger (1964) – as Salvinia natans. Voucher: CHR 234901, 1961. Recognition: Salvinia ×molesta is easily recognised by its floating habit, its dimorphic leaves arranged in whorls of three, and the presence of egg-whisk-shaped hairs on the leaves for buoyancy. When mature and opened out, the floating leaves are shaped like the wings of a butterfly. The plant is sterile, with sporocarps Fig. 10: distribution containing only empty sporangia, or aborted spores, and is probably map based on databased records at of hybrid origin. AK, CHR and WELT. Notes: Salvinia ×molesta has been declared a noxious weed in New Zealand. It is believed to be a hybrid of horticultural origin from

7 South America, now widely distributed in Africa and other areas outside the neotropics (Mitchell 1972). It is a pentaploid with 45 chromosomes (Schneller 1981). It is very similar to S. biloba (syn. S. herzogii) from South America (de la Sota 1962, 1995; Tryon & Tryon 1982). At least some forms of this latter species are also probably of hybrid origin (Schneller 1990b), and are distinguishable from S. ×molesta only by their chromosome number (Schneller 1980). However, there are no records of S. biloba in horticultural cultivation, and infestations outside South America all seem to be referable to S. ×molesta. It has been proposed that the earlier name, S. adnata Desv., should replace S. ×molesta (de la Sota 1995). However, Moran & Smith (1999) argued that S. adnata is of uncertain application, possibly attributable to either S. biloba or S. ×molesta. Desvaux’s type specimen is vegetative, and although de la Sota (2001) provided some characters for distinguishing S. biloba and S. ×molesta, he was unable to examine them in the type specimen itself. The name S. ×molesta should therefore continue to be used for the invasive plant of the Old World tropics.

Fig. 11: Salvinia ×molesta: plant showing tightly Fig. 12: Salvinia ×molesta: plant showing overlapping floating leaves with specialised dimorphic leaves arranged in a whorl of three, with water-repellent hairs on upper surface. two floating green foliar leaves, and one submerged root-like leaf.

Fig. 13: Salvinia ×molesta: a single, Fig. 14: Salvinia ×molesta: plant showing fertile conduplicately folded, foliar leaf with water- axes amongst the submerged leaves, each repellent hairs on upper surface. bearing up to 20 pairs of sporocarps.

8 Fig. 15: Salvinia ×molesta: a single fertile axis Fig. 16: Salvinia ×molesta: water-repellent hairs on amongst the submerged leaves bearing pairs of the upper surface of a foliar leaf, each comprising sporocarps. a papilla with four uniseriate hairs united at their distal ends (shaped like an egg-whisk).

Fig. 17: Salvinia ×molesta: empty microsporangia released from the sporocarps.

9 References

Allan, H.H. 1961: Flora of New Zealand. Vol. I. Indigenous Tracheophyta: Psilopsida, Lycopsida, Filicopsida, Gymnospermae, Dicotyledones. Government Printer, Wellington. Armiger, L.C. 1964: A guide to the more common fresh water plants of the Auckland Province. Tane 10: 49–57. Brown, R. 1810: Prodromus Florae Novae Hollandiae et Insulae Van-Diemen. Johnson, London. Brownsey, P.J.; Given, D.R.; Lovis, J.D. 1985: A revised classification of New Zealand with a synonymic checklist of species. New Zealand Journal of Botany 23(3): 431–489. Brownsey, P.J.; Perrie, L.R. 2013: Azolla rubra revisited. New Zealand Botanical Society Newsletter 111: 6-7. Brownsey, P.J.; Perrie, L.R. 2014: Taxonomic notes on the New Zealand flora: lectotypes in Marsileaceae and Salviniaceae (water ferns). New Zealand Journal of Botany 52(2): 270–271. Brownsey, P.J.; Smith-Dodsworth, J.C. 2000: New Zealand ferns and allied plants. Edition 2. David Bateman, Auckland. Chinnock, R.J. 1998: Azollaceae. In: . Vol. 48. 174–177. Christenhusz, M.J.M.; Zhang, X.-C.; Schneider, H. 2011: A linear sequence of extant families and genera of lycophytes and ferns. Phytotaxa 19: 7–54. de la Sota, E.R. 1962: Contribución al conocimiento de las Salviniaceae neotropicales, III Salvinia herzogii nov. sp. Darwiniana 12: 499–513. de la Sota, E.R. 1995: Nuevos sinónimos en Salvinia Ség. (Salviniaceae, Pteridophyta). Darwiniana 33: 309–313. de la Sota, E.R. 2001: Sobre el tipo de Salvinia adnata (Salviniaceae, Pteridophyta). Boletín de la Sociedad Argentina de Botánica 36: 125–129. Jones, D.L. 1998: Marsileaceae. In: Flora of Australia. Vol. 48. 166–173. Lamarck, J.B.A.P.M. de 1783: Encyclopédie Méthodique. Botanique. Vol. 1. Panckoucke, Paris. Martinov, I.I. 1820: Tekhno-Botanicheskīĭ Slovar'. St Petersburg. Mason, R. 1964: Aquatic weed identification in New Zealand. Proceedings of the 17th New Zealand Weed Control Conference 17: 229–250. Metzgar, J.S.; Schneider, H.; Pryer, K.M. 2007: Phylogeny and divergence time estimates for the fern genus Azolla (Salviniaceae). International Journal of Plant Sciences 168: 1045–1053. Mitchell, D.S. 1972: The Kariba weed: Salvinia molesta. British Fern Gazette 10: 251–252. Mitchell, D.S.; Thomas, P.A. 1972: Ecology of waterweeds in the Neotropics. UNESCO Technical Papers in Hydrology. Vol. 12. UNESCO, Paris. Moran, R.C.; Smith, A.R. 1999: Salvinia adnata Desv. versus S. molesta D.S.Mitch. American Fern Journal 89(4): 268–269. Nagalingum, N.S.; Nowak, M.D.; Pryer, K.M. 2008: Assessing phylogenetic relationships in extant heterosporous ferns (Salviniales), with a focus on and Salvinia. Botanical Journal of the Linnean Society 157: 673–685. Nagalingum, N.S.; Schneider, H.; Pryer, K.M. 2006: Comparative morphology of reproductive structures in heterosporous water ferns and a re-evaluation of the sporocarp. International Journal of Plant Sciences 167: 805–815. New Zealand Weed and Pest Control Society 1969: Standard common names for weeds in New Zealand. Editorial Services Ltd, Wellington. Pryer, K.M. 1999: Phylogeny of marsileaceous ferns and relationship of the fossil Hydropteris pinnata reconsidered. International Journal of Plant Sciences 160: 931–954. Pryer, K.M.; Schuettpelz, E.; Wolf, P.G.; Schneider, H.; Smith, A.R.; Cranfill, R. 2004: Phylogeny and evolution of ferns (monilophytes) with a focus on the early leptosporangiate divergences. American Journal of Botany 91: 1582–1598. Reid, J.D.; Plunkett, G.M.; Peters, G.A. 2006: Phylogenetic relationships in the heterosporous fern genus Azolla (Azollaceae) based on DNA sequence data from three non-coding regions. International Journal of Plant Sciences 167: 529–538. Schneider, H.; Pryer, K.M. 2002: Structure and function of spores in the aquatic heterosporous fern family Marsileaceae. International Journal of Plant Sciences 163: 485–505.

10 Schneller, J.J. 1980: Cytotaxonomic investigations of Salvinia herzogii de la Sota. Aquatic Botany 9: 279–283. Schneller, J.J. 1981: Chromosome numbers and spores of Aublet s.str. Aquatic Botany 10: 81–84. Schneller, J.J. 1990a: Azollaceae. In: Kramer, K.U.; Green, P.S. Pteridophytes and gymnosperms. Vol. 1. In: Kubitzki, K. (ed.) The Families and Genera of Vascular Plants. Springer-Verlag, Berlin. Schneller, J.J. 1990b: Salviniaceae. In: Kramer, K.U.; Green, P.S. Pteridophytes and gymnosperms. Vol. 1. In: Kubitzki, K. (ed.) The Families and Genera of Vascular Plants. Springer-Verlag, Berlin. Séguier, J.F. 1754: Plantae Veronenses. Vol. 3. Verona. Smith, A.R.; Pryer, K.M.; Schuettpelz, E.; Korall, P.; Schneider, H.; Wolf, P.G. 2006: A classification for extant ferns. Taxon 55(3): 705–731. Strasburger, E. 1873: Über Azolla. Dabis, Jena. Svenson, H.K. 1944: The New World species of Azolla. American Fern Journal 34: 69–84. Tindale, M.D.; Roy, S.K. 2002: A cytotaxonomic survey of the Pteridophyta of Australia. Australian Systematic Botany 15: 839–937. Tryon, A.F.; Lugardon, B. 1991: Spores of the Pteridophyta. Springer-Verlag, New York. Tryon, R.M.; Tryon, A.F. 1982: Ferns and allied plants. Springer-Verlag, New York. Wettstein von Westersheim, R. 1903: Handbuch der Systematischen Botanik. Vol. 2. Part 1. Franz Deuticke, Leipzig and Wien.

11 Acknowledgements

We thank the staff at AK, CHR and WELT for loans of specimens and for databasing and providing spreadsheets of collection data. We are grateful to staff at CHR for the preparation of maps and for assistance in editing and formatting the text, and to Nathalie Nagalingum for reviewing the manuscript. P.J. Brownsey and L.R. Perrie Museum of New Zealand Te Papa Tongarewa, PO Box 467, Wellington 6140, New Zealand [email protected] [email protected]

12 Map 1: Map of New Zealand and offshore islands showing Ecological Provinces Map 2: Map of New Zealand showing Ecological Provinces Index

Page numbers are in bold for the main entry, and italic for synonyms. Azolla Lam. 1, 2, 2 Azolla filiculoides var. rubra (R.Br.) Strasb. 5 Azolla pinnata R.Br. 1, 3, 5 Azolla rubra R.Br. 1, 4, 5 Azollaceae Wettst. 2 Salvinia Ség. 1, 2, 6 Salvinia ×molesta D.S.Mitch. 1, 7, 7 Salviniaceae Martinov 1, 2

15 Image Information

Image Creator Copyright License Front cover P.J. Brownsey © Pat Brownsey 1975 CC-BY-NC 3.0 NZ Fig. 1 K. Boardman © Landcare Research 2014 CC-BY 3.0 NZ Fig. 2 L.R. Perrie © Leon Perrie 2013 CC-BY-NC 3.0 NZ Fig. 3 L.R. Perrie © Leon Perrie 2012 CC-BY-NC 3.0 NZ Fig. 4 L.R. Perrie © Leon Perrie 2012 CC-BY-NC 3.0 NZ Fig. 5 K. Boardman © Landcare Research 2014 CC-BY 3.0 NZ Fig. 6 P.J. Brownsey © Pat Brownsey 1975 CC-BY-NC 3.0 NZ Fig. 7 P.J. Brownsey © Pat Brownsey 1975 CC-BY-NC 3.0 NZ Fig. 8 P.J. Brownsey © Pat Brownsey 1975 CC-BY-NC 3.0 NZ Fig. 9 L.R. Perrie © Te Papa 2011 CC-BY-NC 3.0 NZ Fig. 10 K. Boardman © Landcare Research 2014 CC-BY 3.0 NZ Fig. 11 L.R. Perrie © Te Papa 2012 CC-BY-NC 3.0 NZ Fig. 12 L.R. Perrie © Te Papa 2012 CC-BY-NC 3.0 NZ Fig. 13 L.R. Perrie © Te Papa 2012 CC-BY-NC 3.0 NZ Fig. 14 L.R. Perrie © Te Papa 2012 CC-BY-NC 3.0 NZ Fig. 15 L.R. Perrie © Te Papa 2012 CC-BY-NC 3.0 NZ Fig. 16 L.R. Perrie © Te Papa 2012 CC-BY-NC 3.0 NZ Fig. 17 L.R. Perrie © Te Papa 2012 CC-BY-NC 3.0 NZ Map 1 A.D. Wilton © Landcare Research 2014 CC-BY 3.0 NZ Map 2 A.D. Wilton © Landcare Research 2014 CC-BY 3.0 NZ

16 Flora of New Zealand: PDF publications

The electronic Flora of New Zealand (eFloraNZ) project provides dynamic, continually updated, online taxonomic information about the New Zealand flora. Collaborators in the project are Landcare Research, the Museum of New Zealand Te Papa Tongarewa, and the National Institute of Water and Atmospheric Research (NIWA). The eFloraNZ presents new systematic research and brings together information from the Landcare Research network of databases and online resources. New taxonomic treatments are published as fascicles in PDF format and provide the basis for other eFloraNZ products, including the web profiles. eFloraNZ will have separate sets of PDF publications for , lichens, liverworts and hornworts, mosses, ferns and lycophytes, and seed plants. For each eFloraNZ set, the PDF files are made available as dated and numbered fascicles. With the advent of new discoveries and research, the fascicles may be revised, with the new fascicle being treated as a separate version under the same number. However, superseded accounts will remain available on the eFlora website.

Fern and Lycophyte Set (ISBN 978-0-478-34761-6) The Fern and Lycophyte Set includes ferns and lycophytes indigenous to New Zealand, together with exotic species that have established in the wild. Species that are found only in cultivation are excluded. Editor-in-Chief: Ilse Breitwieser Series Editors: Ilse Breitwieser (Principal), Peter Heenan, Aaron Wilton Steering committee: Ilse Breitwieser, Pat Brownsey, Peter Heenan, Wendy Nelson, Aaron Wilton Technical production: Aaron Wilton with Kate Boardman, Bavo de Pauw, Sue Gibb, Ines Schönberger, Katarina Tawiri, Margaret Watts Copy Editor: Christine Bezar

17 ISBN 978-0-478-34779-1

9 780478 347791